The individual and social drivers of primate innovation

The individual and social drivers of primate innovation

How does nonhuman primate innovation compare to our own? Many primates innovate, for

example to get otherwise inaccessible food or to increase their social standing, and

nonhuman primate innovation can be broken into three component steps. It begins with

the initial invention, which is then transmitted to other members of the inventor’s group,

and is then adopted by other individuals and maintained within the society. These three

steps – invention, transmission, and maintenance – are all required for innovation and in

this review, I discuss the factors (social, environmental, and cognitive) that influence each

step. I also highlight the comparable and contrasting features between human and

nonhuman primate innovation. In contrast to human innovations, primate innovations are

relatively simple and are typically self-serving. Nonhuman primates do not invent new

products explicitly for the use of others (although group members certainly copy others'

innovations) and nor are their inventions artistic or abstract in nature. Intriguingly,

although chimpanzees and other nonhuman primates appear to be expert at copying

others’ inventions, there is far less evidence of their ability to build upon others’ inventions

(i.e., to show cumulative culture). At the core of our complex cultural world is the fidelity

with which we copy others and our specialism at building upon the ideas of others. Thus, it

is the cumulative nature of our innovative process that has created our complex material

cultural world and is a key difference between how we innovate, learn and transmit

knowledge, and how our chimpanzee cousins copy one another. Another difference is our

ability to work collaboratively in teams to innovate and develop new technologies, as well

as our potential to cooperate in an altruistic way that allows for planning for future

generations. In conclusion, perhaps primate innovation can be most usefully likened to

human ‘user innovators’ who typically innovate products or techniques to fill a personal

need, rather than by being driven to create a product to go to market.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2276v1 | CC BY 4.0 Open Access | rec: 11 Jul 2016, publ: 11 Jul 2016

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The Individual and Social Drivers of Primate Innovation 1

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Lydia M Hopper 3

Lester E. Fisher Center for the Study and Conservation of Apes, Lincoln Park Zoo, Chicago, IL 60614 4

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WHAT IS (NONHUMAN PRIMATE) INNOVATION? 6

Innovation is not a solitary pursuit. Inventors are shaped by their social environment and rely on their 7 community to adopt their inventions. Furthermore, innovation is not a one-way process in which users 8 only adopt others’ innovations, but users themselves may also be active in product development and 9 modification (von Hippel, 2005). In turn, discoveries are amended, improved upon, or discarded, 10 streamlining and refining innovations in an iterative fashion. This ratcheting creates an accumulation of 11 cultural complexity (Tomasello et al., 1993), a process which Matt Ridley famously crystalized and 12 popularized as “ideas having sex” (Ridley, 2010). There is a feedback loop within the social environment 13 as individuals innovate through trial-and-error and also by copying and improving upon the ideas of 14 others. The importance of social information for how we learn and innovation has long been recognized. 15 For example, in her book Openness, Secrecy, Authorship, Pamela Long (2001) noted that Vitruvius 16 believed that humans discovered the art of building in part by imitating the nests of swallows1. Thus, 17 innovations can arise both through personal discovery and also by adapting the ideas and behavior of 18 others (in this case, replicating and expanding upon the nest making of birds2), but what drives our need 19 to innovate? 20

We can innovate in a methodical planned way, in an attempt to address a current need, or we might 21 innovate spontaneously, without forethought or clear understanding of our goal. The former relies on 22 spotting gaps in the market or in our needs, and potentially the ability for mental time travel: future 23 planning that allows us to predict and prepare for future events or needs (Vale et al., 2012). The latter 24 arises from simple trial-and-error problem solving – such inventions are serendipitous and do not rely on 25 cognitive planning or forethought. How much intentionality and active learning is involved in innovation 26 has been much debated and some scholars divide innovation into active versus passive innovation (also 27 referred to as Type I versus Type II innovation) (Reader et al., 2016). Simply put, active innovation 28 requires learning and insight, while passive innovation can arise more serendipitously. Beyond our 29 underlying understanding and motivation to innovate, the process of innovation has been 30 subcategorized as (1) invention, (2) refinement, (3) recombination, and (4) exaptation (Mesoudi et al., 31 2013). In this way, an invention can be more than a new product or solution, it can also be the 32 application of an already-existing invention repurposed in a new way, the combination of new existing 33

1 The parallels between birds’ nests and our abodes was also a topic of interest for Vincent van Gogh who noted in in text accompanying a sketch he drew of a bird’s nest that he sent to his brother Theo “The nestlings and the nests, I feel deeply for them – especially people’s nests, those huts on the heath and their inhabitants.” Text from a letter written by van Gogh on September 4th 1885 translated by Johanna van Gogh-Bonger and reproduced by WebExhibits: http://www.webexhibits.org/vangogh/letter/15/425.htm 2 Pamela Long (2001) also reported “the atomist Democritus of Abdera articulated… *that+ humans discovered arts

such as weaving and building houses by imitating the animals; they invented weaving, for example, after observing how spiders create their webs” (Long, p. 19)


http://www.webexhibits.org/vangogh/letter/15/425.htm

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technologies to create a novel product, or the refinement of an existing product in a way that creates a 34 novel innovation3. 35

It has been proposed that imagination, which is linked to our ability to remember and predict events, is 36 a key component of creativity, itself a potential driver of innovation. Creativity is an intriguing and 37 somewhat elusive property, and there is much individual variation in peoples’ creativity, although 38 current research is helping to elucidate the neural underpinnings of creativity (Bashwiner et al., 2016; 39 Jung et al., 2016). Creativity is inherently playful, and innovations driven by creativity arise from 40 curiosity, not need (van Schaik et al., 2016). Yet the adage states that necessity is the mother of 41 invention (not creativity). I reality, it is likely that, for humans, both need and creativity can drive 42 innovation, although there is likely to be individual and situational variation. Determining whether an 43 individual’s tinkering is driven by curiosity, rather than need, may be difficult, especially for nonverbal 44 nonhuman primates. Considering primates, ‘environmental opportunity’ actually appears to be a better 45 predictor of when monkeys and apes make and use tools (Koops et al., 2014). Essentially, a chimpanzee 46 cannot invent a new kind of stone tool if there are no stones lying around, however badly she needs to 47 crack open a nut, or how curious she is. Furthermore, captive primates are thought to be more 48 innovative than their wild counterparts because they have more ‘spare time’ in which to explore. 49 Kathelijne Koops and colleagues (2014) concluded that when and how wild chimpanzees, orangutans 50 and capuchin monkeys make tools can be explained by a combination of environmental opportunity, 51 cognitive ability, and sociality, all of which differentially influence when and if individuals innovate and 52 when and if others adopt their innovations. It is likely that creativity is not a pre-requisite for innovation, 53 especially if we consider the innovations of nonhuman primates. 54

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INNOVATION BY NONHUMAN PRIMATES 56

A classic case of primate innovation is that of Imo, the Japanese macaque, who discovered that if she 57 washed sand-covered potatoes left on the beaches of Koshima Island, Japan, by researchers trying to 58 study her group’s behavior, they would be more palatable (Yamagiwa, 2010). Primatologists in the late 59 1940s and early 1950s studying the Japanese macaques that lived on Koshima Island placed potatoes on 60 the sea shore to lure the monkeys out of the forests where they lived, making it easier for the 61 researchers to observe the monkeys’ behavior4. Imo (whose name means potato in Japanese) was the 62 first to wash her potatoes in a nearby freshwater stream. Later, she innovated for a second time and 63 carried the potatoes to the sea to wash off the sand before eating them; not only did washing the 64 potato in the sea remove the sand, but this latter variant added a salty seasoning to the food. Imo was 65 quickly copied by offspring, and soon the behavior spread throughout the troop (de Waal & Bonnie, 66 2009). Such socially-mediated learning has been dubbed the ‘second inheritance system’ (Whiten, 67 2005): a behavior is spread throughout a group of individuals via observational learning rather than by 68 genetic inheritance. 69 3 The adaptation of previously-made inventions for new markets or cultures is a process that is alive and well in current day China. As Clive Thompson (2015) noted in his article about Chinese “copy cat” firms, local firms in China had an edge over the established giants from abroad because they had local cultural understanding and so were able to adapt existing technologies to better suit the local market. The examples Thompson provides demonstrate that not only can replications be considered innovations when applied in a new setting, but that they also spawned future (novel) innovations. 4 The Japanese primatologists studying Imo and her troop were also innovative, as unlike many behaviorists before

them who studied animals at the population or species level, Kinji Imanishi, Shunzo Kawamura, and Jun’ichiro Itani were interested in studying animal societies at the level of unique and identifiable individuals (Hirata et al., 2001).


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Imo’s potato-washing innovation is a perfect example of what is considered as innovation by 70 primatologists: a novel behavior, invented by a single individual, is adopted by others in the innovator’s 71 group (Laland & Reader, 2003). Of note too, and in contrast to human innovations, Imo’s innovation was 72 relatively simple and was very self-serving. She was not inventing a new product for the use of others in 73 her group (although they certainly exploited her innovation by copying her) and nor was her invention 74 artistic or abstract in nature. We are unique in the scale and scope of our innovations, even as compared 75 to our closest living cousins, the chimpanzees. Chimpanzees are renowned for their cognitive prowess, 76 and are both expert innovators and keen observers of their group mates’ behavior allowing them to 77 copy the innovations of others. They have been documented to create and use a variety of tools, which 78 appear to vary culturally across different societies, and to develop novel behavioral and gestural cultures 79 (Whiten et al., 1999). Despite this, they do not have the complexity of material culture that we do; they 80 do not live in cities, communicate over long distances using technologies designed for that purpose, nor 81 do they use symbolic representation (Hill, 2009). 82

When we consider primate innovation, creativity and intention are not always inferred, while when we 83 discuss human innovation, we often consider innovation to be insightful – entrepreneurs aim to spot 84 gaps in the market and launch the product to market. Primates, by contrast, are typically not trying to 85 innovate to create a product, tool, or skill that others will adopt. They do not teach, and any 86 transmission of information is passive (Matsuzawa et al., 2001). Given this, perhaps primate innovation 87 can be more usefully likened to ‘user innovators.’ Eric von Hippel (2005) provides a comprehensive 88 overview of the current activity and importance of user innovators who typically innovate products or 89 techniques to fill a personal need, rather than by being driven to create a product to go to market. For 90 example, in addition to innovating and creating products for sale, manufacturing companies invent and 91 develop tools for their own use, that enhance the fabrication process of their products. These user 92 innovations can be in the form of modifications to existing tools or the invention of a completely novel 93 tool. Primates, like user innovators, typically create tools for their own needs or adapt the tools used by 94 others. In both cases, their inventions are for use by them personally, rather than to distribute within 95 their community. Any distribution that is observed, would be unintentional on the part of the inventor, 96 and happen passively by social learning as shown experimentally in a number of experiments (Hopper, 97 in press, provides a review). 98

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THE PROCESS OF INNOVATION 100

As highlighted through the example of Japanese monkey Imo’s potato washing described above, 101 innovation is process that can be broken in three component steps. It begins with the initial invention, 102 which is then transmitted to other members of the inventor’s group, and is then adopted by other 103 individuals and maintained within the society. These three steps – invention, transmission, and 104 maintenance – are all required for innovation and yet the factors that influence each step vary (Brosnan 105 & Hopper, 2014 provide a review of the mechanisms that can inhibit each of these three steps when 106 animals innovate). 107

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Inventive Inventors 109

“In order to educate and support innovative leaders, we should first identify what characterizes them” 110 Rebecca Bagley wrote in a 2014 Forbes article entitled The 10 Traits of Great Innovators, before going on 111


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to list ten characteristics of innovative personalities. Such an approach is not uncommon and highlights 112 both our desire to pinpoint the specific characteristics of innovators and also the inherent difficulties in 113 doing so. It is clear that there is no single trait associated with ‘innovativeness’ in humans, and the same 114 appears to be true for nonhuman primates and other animals. 115

In a recent study run with a group of chimpanzees housed at Chicago’s Lincoln Park Zoo, the 116 chimpanzees were provided with plastic tokens that they could exchange with researchers who stood at 117 the perimeter of their enclosure (Hopper, Kurtycz et al., 2015). The experiment was designed such that if 118 the chimpanzees took their tokens to the experimenter who was standing furthest away and exchanged 119 it with them, they received a highly-desirable food reward (for them, this was a grape). Contrastingly, if 120 the chimpanzees choose to carry their tokens to the nearer researcher they received a less-desirable 121 piece of carrot for each token exchanged. Importantly, the chimpanzees were not trained how to 122 exchange tokens with the researchers, and nor were they trained about the relative value of the food 123 rewards available at each location. The beta-ranking male chimpanzee, Optimus Prime, was the first 124 member of the group to ever exchange a token for a food reward and he did so with the researcher 125 standing closest by, gaining a piece of carrot. The rest of his group then quickly learned his new skill, all 126 exchanging their tokens for the readily-accessible, but less-desirable, carrot pieces. The seventh time 127 that the chimpanzees were presented with tokens, the lowest-ranked member of the group, a 13-year 128 old female named Chuckie innovated. She was the first in her group to discover that if she carried her 129 tokens a little further, she could exchange them for better rewards. She was also the first to do so again 130 in a later phase of the study when the locations where the researchers stood to exchange tokens 131 changed. What was it about Chuckie and Optimus Prime that encouraged them to innovate how and 132 where to exchange the otherwise seemingly-useless plastic tokens? A review of innovations by wild 133 primates revealed that chimpanzees’ innovative tendencies are influenced by their age, rank and sex, 134 with juvenile, low-ranking, or male chimpanzees being more likely to innovate, most likely to enable 135 them to secure otherwise inaccessible resources (Reader & Laland, 2001). This pattern mirrors that seen 136 among the chimpanzees at Lincoln Park Zoo. The first to innovate, Optimus Prime, was lower-ranking 137 and male and, Chuckie was very low ranking young member of the group. 138

More recent studies with captive chimpanzees investigating how chimpanzee personality traits correlate 139 with their problem-solving prowess have reported that chimpanzees rated highly on personality factors 140 related to curiosity, exploration, and persistence are more dogged in their efforts to solve the puzzles 141 presented to them and are ultimately more successful in solving them (Massen et al., 2013; Hopper et 142 al., 2014). Ultimately, it is most likely that an individual’s innovative tendencies arise from a combination 143 of factors. Therefore, ‘innovativeness’ can likely be considered as an emergent property comprising of 144 internal states and predispositions as well as external environmental factors (Reader et al., 2016). 145 Beyond simply considering cognitive skills, it is also important to consider motor flexibility, learning and 146 physiology, all of which influence the different ways in which animals can interact with their physical 147 environment and, therefore, what and how they innovate (Griffin, 2016). 148

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Transmission and Adoption 150

We can all call to mind specific inventors, individual’s whose tireless work or genius insight resulted in 151 the creation of a novel creation: Henry Ford who founded the Ford Motor Company, Sir Timothy John 152 Berners-Lee who invented the World Wide Web, and Ada Lovelace who is recognized as the first 153 computer programmer when she wrote her notes on algorithms to be computed by machines (Isaacson, 154 2014). However, even the most highly inventive individuals are inspired by those around them (whether 155


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directly or not). So, beyond the intrinsic factors that some individuals have that drive them to innovate, 156 it is absurd to think that individuals are not influenced by the society around them. As Muthukrishna 157 and Henrich (2016) noted “innovations arise as an emergent consequence of our species’ psychology 158 applied within our societies and social networks” and there is a swathe of research showing how, just 159 like us, nonhuman primate decision making is also influenced by their social environment (Cronin & 160 Hopper, in press; Hopper & Cronin, in press). Beyond this, the social environment in which an innovator 161 lives is key as it is what allows the innovation to be propagated and distributed (Rogers, 2003). 162

Nonhuman primates are more explorative and less fearful to approach new things (neophobic) when 163 they have social support. Studies with captive New World monkeys, including capuchin monkeys and 164 squirrel monkeys, have shown that they are much more likely to solve novel puzzles when they are 165 tested with their group mates present, than when tested alone (Dindo et al., 2009). For example, 166 squirrel monkeys that were given a box that contained a meal worm (a tasty treat for a squirrel 167 monkey!) were only able to learn how to slide open a door on the box to retrieve the meal worm if they 168 had a companion in the testing booth with them, whereas those monkeys tested by themselves never 169 solved the task despite its relative simplicity (Hopper et al., 2013). 170

Beyond providing social support and encouraging exploration, being in the presence of group mates 171 allows primates to exploit the behavior of others and replicate their actions and innovations, which 172 saves them the potential costs associated with trial-and-error learning (Hopper, in press). For example, 173 experimental research with captive and wild primates has shown that they can socially learn how to 174 assemble and use tools from observing others (Price et al., 2009), how to solve problems presented on 175 touchscreen computers (Subiaul et al., 2004), and which foods to eat or avoid, even if that contradicts 176 their own personal experiences (van de Waal et al., 2013). Social learning is important because it is the 177 key mechanism that facilitates transmission of innovations within social groups – primates are experts at 178 gaining new skills this way – and it is the first step of cultural diversification. Via social learning, primates 179 learn skills that will sustain them, such as how to capture and process prey or how to make and use 180 tools, as well as social gestures and customs, which are important for maintaining social bonds (Hopper 181 et al., 2011). Interestingly, just as certain individual characteristics are associated with greater 182 innovativeness, certain primates are more likely to copy the behavior of others, which is influenced by 183 factors including an individual’s personality (Carter et al., 2011) and rank (Kendal et al., 2015), as well as 184 their species (Pasquaretta et al., 2014) (Hopper, in press, provides a review). 185

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Imitating and Improving Others’ Inventions 187

Perhaps more importantly than simply ‘blindly’ copying the behaviors of others, is the ability to build 188 upon the ideas of others. This ‘ratcheting’ effect (Tomasello et al., 1993) is a keystone of human culture. 189 A key characteristic of cumulative cultural artifacts is that contemporary inventions could not be 190 invented by a single individual; they are the creation of multiple generations’ tinkering and modifying. 191 Cumulative culture further emphasizes the idea that innovators are not uninfluenced by their social 192 environment, whether their current social group or previous generations. We are all standing on the 193 shoulders of giants. Intriguingly, although chimpanzees and other nonhuman primates appear to be 194 expert imitators (apes really do ape), there is far less evidence of their ability to build upon others’ 195 inventions (i.e., to show cumulative culture, Mashall-Pescini & Whiten, 2008). While some groups of 196 wild chimpanzees have an array of over 20 different tools in their tool kit, in only a few circumstances do 197 they use different tools in combination (so-called ‘tool sets,’ e.g., Sanz, Schoning & Morgan 2009). 198 Furthermore, chimpanzees have never been observed to combine tools, to add on elements to tools 199


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(such as adding a handle), or to create tools to make other tools. Thus, although chimpanzees use tools 200 that are differentiated, and show forethought in their creation and use of tools (Sanz, Call & Morgan, 201 2009; Hopper, Tennie et al., 2015), all their tools could likely be invented by a single individual, and 202 cannot be considered as the result of an accumulation of techniques (Tennie et al., 2009). 203

What is it about our ability not just to copy others, but to extend upon their inventions, that 204 chimpanzees lack? Lewis Dean and colleagues (2014) reviewed characteristics that might be unique to 205 us and explain why we, but not chimpanzees, evidence complex cumulative culture. They cited cognitive 206 skills including our ability to innovate and to faithfully imitate the behavior of others; our propensity to 207 work collaboratively and to share; and that we actively teach one another and can communicate 208 complex instructions and descriptions. As the idea of a single innovative trait seems too reductionist, so 209 does the idea that a single characteristic is what differentiates our social learning skills from our 210 chimpanzee cousins. It is likely that multiple cognitive, social and ecological factors inhibit chimpanzees’ 211 ability or drive to develop complex cultural artifacts, tools and customs. Contemporary experimental 212 research (e.g., Dean et al., 2012) and theoretical research (e.g., Lewis & Laland, 2012) is now just 213 beginning to tap into the mechanisms that promote and hinder cumulative cultural transmission in us 214 and other species (Caldwell et al., 2012). 215

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SOCIAL NETWORKS, DOMINANCE, AND FRIENDSHIPS 217

Although chimpanzees appear less likely than us to (intentionally) modify and improve upon previous 218 generations’ innovations, they are certainly skilled at copying the actions of others and replicating their 219 inventions. In this way, inventions can become spread within a community. However, the likelihood that 220 an invention will be adopted by the innovator’s social group and spread widely within their community 221 is dependent on the place of the innovator in their social network, and the dynamics of their network. 222 From 40 years of observing wild chimpanzees living in Tanzania, Toshisada Nishida and colleagues (2009) 223 concluded that “innovation was not rare, but the emergence of fashion or establishment of traditions 224 seems to occur rarely in chimpanzee society.” This nicely highlights the important interplay between the 225 individual innovator and their society with regard to the adoption and transmission of inventions. When 226 considering the inventions of human entrepreneurs, we often merit the success of an invention as to 227 whether it reached commercialization, which may be related to the relevance or ‘excellence’ of the 228 invention itself (Scott et al., 2015) or how well connected the entrepreneur is (Lee, 2015). While it may 229 be a complex exercise to evaluate the merit of chimpanzees’ innovations, through the use of social 230 network analysis, combined with observational and experimental research focused on the transmission 231 of inventions, it is possible to dissect how the social dynamics of a social group might influence the likely 232 transmission of an invention (Hobaiter et al., 2014) and the stability of the social system (Fushing et al., 233 2014). 234

In her study of human innovation transmission and the characteristics of communities that foster 235 entrepreneurship, Minha Lee (2015) identified three key aspects of human societies that would promote 236 knowledge dissemination and the transfer of inventions. The first two related to defining how 237 interconnected the social group is; the number of individuals who engage positively with one another 238 (‘density’) and the number of connections that key individuals have (‘central connectors’). The third 239 component was the relative knowledge that individuals within a group have, with those individuals with 240 key knowledge (‘knowledge bases’) and their connectivity with central connectors being key to enable 241 innovation. Similarly, experimental work with captive primates (e.g., squirrel monkeys: Claidière et al., 242 2013) and observations of wild groups (e.g., chimpanzees: Hobaiter et al., 2014) have revealed that an 243


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individual’s social relationship with an inventor predicts their likelihood of copying that inventor and 244 adopting their invention; birds of a feather flock together, and then go on to copy one another. Nicolas 245 Claidière and colleagues (2013) also found that more centrally-connected squirrel monkeys were more 246 likely to learn new innovations and would also adopt the innovation more quickly than other members 247 of the group, as has also been shown within wild communities of birds (Alpin et al., 2015). 248

Beyond the number of social connections a primate has, the quality of those relationships is also 249 important, something that varies both within, as well as across, species (Cronin et al., 2014; Pasquaretta 250 et al., 2014). Many primate societies are governed by strong dominance hierarchies, and, as discussed 251 above, low-ranking chimpanzees are more likely to innovate than dominants, but they are also more 252 likely to copy the behavior of others than are dominants (Kendall et al., 2015). The role of subordinate 253 chimpanzees in the innovative process (i.e. their propensity to innovate and also to copy others) may 254 also explain Nishida and colleagues’ (2009) report that many wild chimpanzee inventions were not 255 adopted by their group mates5. Chimpanzees typically only look to older and more dominant individuals 256 for information, and are less likely to copy individuals that are subordinate or younger individuals (Biro 257 et al., 2003). This applies to low-ranking individuals and juveniles within a group, and also chimpanzees 258 that are low-ranking because they immigrated into a new group. The low rank of immigrant (typically 259 female6) chimpanzees also inhibits transmission of information between chimpanzee communities, as 260 well as within them, as residents are less likely to copy to the behavior of immigrants, even if they arrive 261 with novel skills (Matsuzawa & Yamakoshi, 1996; Biro et al., 2003). 262

Wild chimpanzees that join new communities typical adopt the traditions of the new group; they 263 conform to the majority. Recent observations of two communities of chimpanzees in Côte d’Ivoire 264 revealed the perhaps surprising insight that when females left their natal group, in which they had a 265 culture of using wooden tools to crack open nuts, and joined the neighboring troop that typically used 266 stone tools to crack nuts, within a couple of months, the immigrant females too were predominantly 267 using stone tools to crack nuts (Luncz & Boesch, 2014). This transition from wooden to stone tools arose 268 despite the females already knowing an equally efficient strategy, and one which they had used all their 269 life previously. Just as chimpanzees typically copy the majority, they also ignore the minority. This is 270 highlighted by the example described by Matsuzawa and Yamakoshi (1996) of a female chimpanzee in 271 Guinea, called Yo, who was an immigrant to her group. The Japanese researchers provisioned the 272 chimpanzee group with novel coula nuts, which are hard shelled and difficult to crack open. Almost 273 immediately Yo placed a coula nut on a stone ‘anvil’ and used a second stone hammer to crack open the 274 nut. It is probable that Yo had experience cracking these nuts in her natal group and so likely did not 275 invent this behavior. What is striking though, is that despite ‘demonstrating’ this neat new trick to her 276 group, the only other chimpanzees to follow her lead and start cracking open and eating the coula nuts 277 were two youngsters. None of the adults copied and the behavior never spread within the group. 278

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5 This interplay between a chimpanzee’s individual status and innovation (invention and transmission) is reminiscent of the incumbent’s curse, which posits that large dominant firms are less nimble than smaller companies and startups and are less likely to innovate, or, if they do, they do so slowly in small incremental steps. However, and as with innovation by dominant chimpanzees, the lore that large and old firms is not always borne out (Chandy & Tellis, 2000). 6 Female chimpanzees typically leave their natal group around adolescence and join neighboring communities; an evolutionary strategy for avoiding inbreeding.


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CONCLUDING THOUGHTS 281

We are a species renowned for our innovative abilities, both to invent independently and to copy the 282 innovations of others. At the core of our complex cultural world is the fidelity with which we copy 283 others7, allowing for faithful transmission and adoption of innovations within a community, and also our 284 specialism at building upon the ideas of others. Andrea Griffin (2016) characterized this as ‘connective 285 thinking’, while Matt Ridley coined the term ‘ideas having sex.’ Indeed, as Eric von Hippel (2005) noted, 286 “to say an innovation is minor is not the same as saying it is trivial: minor innovations are cumulatively 287 responsible for much or most technical progress” (p. 21). Thus, it is the cumulative nature of our 288 innovative process that has created our complex material cultural world and is a key differentiator 289 between how we innovate, learn and transmit knowledge, and how our chimpanzee cousins copy one 290 another (Tennie et al., 2009; Dean et al., 2014). 291

Another difference is our ability to work collaboratively in teams to innovate and develop new 292 technologies, as well as our potential to cooperate in an altruistic way that allows for planning for future 293 generations (Stout, 2015). While chimpanzees can cooperate, for example when hunting, it does not 294 appear that they collaborate in the process of innovating as is seen among human teams (Cronin & 295 Hopper, in press). With our communicative skills, we can transmit ideas via written or oral instruction, 296 whereas primates typically only learn from others via direct observation. Some primate social learning 297 can occur indirectly, for example a chimpanzee might learn how to crack nuts by discovering the 298 discarded hammer and anvil used by a group mate, but this appears to be a less effective and efficient 299 transmission stream (Caldwell et al., 2012). 300

Although nonhuman primates have not created the diversity of material technologies that we have, nor 301 do they build and live in complex cities, they nonetheless are expert innovators. Innovations by primates 302 have been reported in a range of realms including tool construction (e.g., Hobaiter et al., 2014), the 303 eating and processing of novel foods (e.g., Leca et al., 2008) and the invention of novel gestures to 304 communicate meaning and maintain social bonds (e.g., Laidre, 2008). They are also experts in the social 305 realm. Highly intelligent, political, and social creatures that can navigate the complexities of group living 306 through the formation of alliances and friendships. Currently, more than half of the world’s primate 307 species are facing extinction, so perhaps their biggest challenge to date is to innovate and adapt to the 308 changing demands of their fragmented habitat and the anthropogenic pressures they encounter. 309

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ACKNOWLEDGMENTS 311

The ideas presented in this chapter arose from many inspiring discussions at meetings hosted by the 312 Gruter Institute for Law and Behavioral Research. In particular, I wish to thank Monika Gruter Cheney, 313 Oliver Goodenough, Andrew Torrance, Susan Dudley, Lynn Stout, Kevin McCabe and Sean O’Connor for 314 particularly fruitful discussions that helped me explore new perspectives on how chimpanzee innovation 315 compares and contrasts with our own. Thanks too to Andrew Steets and Sarah Jacobson for giving me 316 useful feedback on early drafts of this chapter. I also wish to thank Claudio Tennie, Sarah Brosnan, 317 Emma Flynn, Steve Ross, Andy Whiten, and Rachel Kendal with whom I have studied many of the ideas 318 relating to chimpanzee innovation and social learning discussed herein. At the time of writing, I was 319 supported by the Leo S. Guthman Fund. 320

7 Indeed, developmental psychologist Andy Meltzoff dubbed us Homo imitans (Metlzoff 1998).


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The individual and social drivers of primate innovation

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